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<document ID-DOI="http://doi.org/10.5281/zenodo.6093830" ID-GBIF-Dataset="e7e4f349-2d92-4848-a7a6-e61ce29bd0c7" ID-GBIF-Taxon="114926070" ID-ISSN="1175-5326" ID-Zenodo-Dep="6093830" ID-ZooBank="EDF9E69E-7898-4CF8-B447-EFF646FE3B44" checkinTime="1447312844506" checkinUser="donat" docAuthor="Taylor, Robert W." docDate="2015" docId="232E11751E71FFDFFF085CA9FC4632CC" docLanguage="en" docName="29626.pdf" docOrigin="Zootaxa 3911 (4)" docTitle="Myrmecia impaternata Taylor, 2015, sp.n." docType="treatment" docVersion="5" lastPageId="23" lastPageNumber="516" masterDocId="DF17690D1E64FFC8FF9F5C3EFFA23574" masterDocTitle="Ants with Attitude: Australian Jack-jumpers of the Myrmecia pilosula species complex, with descriptions of four new species (Hymenoptera: Formicidae: Myrmeciinae)" masterLastPageNumber="520" masterPageNumber="493" pageId="21" pageNumber="514" updateTime="1644949943975" updateUser="ExternalLinkService">
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<mods:title>Ants with Attitude: Australian Jack-jumpers of the Myrmecia pilosula species complex, with descriptions of four new species (Hymenoptera: Formicidae: Myrmeciinae)</mods:title>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Taylor, Robert W.</mods:namePart>
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<mods:title>Zootaxa</mods:title>
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<mods:date>2015</mods:date>
<mods:detail type="volume">
<mods:number>3911</mods:number>
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<mods:number>4</mods:number>
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<mods:start>493</mods:start>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.11646/zootaxa.3911.4.2</mods:identifier>
<mods:identifier type="ISSN">1175-5326</mods:identifier>
<mods:identifier type="ZooBank">EDF9E69E-7898-4CF8-B447-EFF646FE3B44</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.6093830" ID-GBIF-Taxon="114926070" ID-Zenodo-Dep="6093830" LSID="urn:lsid:plazi:treatment:232E11751E71FFDFFF085CA9FC4632CC" httpUri="http://treatment.plazi.org/id/232E11751E71FFDFFF085CA9FC4632CC" lastPageId="23" lastPageNumber="516" pageId="21" pageNumber="514">
<subSubSection box="[151,493,151,177]" pageId="21" pageNumber="514" type="nomenclature">
<paragraph box="[151,493,151,177]" pageId="21" pageNumber="514">
<heading bold="true" box="[151,493,151,177]" fontSize="11" level="1" pageId="21" pageNumber="514" reason="1">
<emphasis bold="true" box="[151,493,151,177]" pageId="21" pageNumber="514">
<emphasis bold="true" box="[151,427,151,177]" italics="true" pageId="21" pageNumber="514">
<taxonomicName box="[151,427,151,177]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata" status="sp.n.">Myrmecia impaternata</taxonomicName>
</emphasis>
<taxonomicNameLabel box="[434,493,152,177]" pageId="21" pageNumber="514" rank="species">sp.n.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[151,295,186,211]" pageId="21" pageNumber="514" type="description">
<paragraph box="[151,295,186,211]" pageId="21" pageNumber="514">
(
<figureCitation box="[159,286,186,211]" captionStart="FIGURES 16 18" captionStartId="22.[151,263,1956,1977]" captionTargetBox="[282,1304,202,1933]" captionTargetId="figure@22.[282,1305,193,1933]" captionTargetPageId="22" captionText="FIGURES 16 18. Myrmecia impaternata, National Botanic Gardens, Canberra, ACT. Standard views. HW 2.29 mm, WL 3.35 mm, PW 1.42 mm." pageId="21" pageNumber="514">Figs 1618</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection pageId="21" pageNumber="514" type="discussion">
<paragraph pageId="21" pageNumber="514">
<taxonomicName box="[151,411,259,283]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[151,411,259,283]" italics="true" pageId="21" pageNumber="514">Myrmecia impaternata</emphasis>
</taxonomicName>
is broadly sympatric with
<taxonomicName box="[724,869,258,283]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="croslandi">
<emphasis box="[724,869,258,283]" italics="true" pageId="21" pageNumber="514">M. croslandi</emphasis>
</taxonomicName>
, a matter of possible biological significance (see below). Both species are common in and around Canberra and on the New England Tableland around Armidale. There are confirmed Queensland records of
<taxonomicName box="[648,790,330,355]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="croslandi">
<emphasis box="[648,790,330,355]" italics="true" pageId="21" pageNumber="514">M. croslandi</emphasis>
</taxonomicName>
from the Darling Downs in extreme SE Queensland, and of
<emphasis box="[186,358,367,391]" italics="true" pageId="21" pageNumber="514">
<taxonomicName box="[186,358,367,391]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">M impaternata</taxonomicName>
</emphasis>
from Tamborine Mountain south of Brisbane. JACP voucher specimens of this taxon were discussed by Imai, Taylor
<emphasis box="[456,516,402,427]" italics="true" pageId="21" pageNumber="514">et al.</emphasis>
(1994) as “PBF1 hybrids”, with 2 types: PBF11 and PBF12 (PB =
<taxonomicName box="[1322,1412,402,427]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="pilosula">
<emphasis box="[1322,1412,402,427]" italics="true" pageId="21" pageNumber="514">pilosula</emphasis>
</taxonomicName>
x
<taxonomicName box="[151,224,441,466]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="banksi">
<emphasis box="[151,224,441,466]" italics="true" pageId="21" pageNumber="514">banksi</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection pageId="21" pageNumber="514" type="materials_examined">
<paragraph pageId="21" pageNumber="514">
<emphasis bold="true" box="[199,361,477,501]" pageId="21" pageNumber="514">Type locality.</emphasis>
Immediately E to NE of Corang River Bridge (-35 12, 150 03) on the Nerriga Road, near Braidwood, NSW. This type-locality is common also to
<emphasis box="[818,965,513,538]" italics="true" pageId="21" pageNumber="514">
<taxonomicName box="[818,965,513,538]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="croslandi">M. croslandi</taxonomicName>
</emphasis>
(see above). The site, also supports
<emphasis italics="true" pageId="21" pageNumber="514">
<taxonomicName class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">M. impaternata</taxonomicName>
</emphasis>
,
<emphasis box="[302,444,549,573]" italics="true" pageId="21" pageNumber="514">
<taxonomicName box="[302,444,549,573]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="croslandi">M. croslandi</taxonomicName>
</emphasis>
and
<emphasis box="[498,625,549,573]" italics="true" pageId="21" pageNumber="514">
<taxonomicName box="[498,625,549,573]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="pilosula">M. pilosula</taxonomicName>
</emphasis>
(Eastern Race).
</paragraph>
<paragraph pageId="21" pageNumber="514">
<emphasis bold="true" box="[199,395,585,610]" pageId="21" pageNumber="514">Type deposition.</emphasis>
Holotype and paratypes in ANIC, paratypes or type-compared vouchers in AMSA, MVMA, QMBA, SAMA, WAMA, TMHA) and in BMNH, CASC, MCZC, MHNG..
</paragraph>
<paragraph pageId="21" pageNumber="514">
<emphasis bold="true" box="[199,588,657,682]" pageId="21" pageNumber="514">Material examined, distribution.</emphasis>
QUEENSLAND: Mt Tamborine [-27 55, 153 10], O. Deane (MVMA); Mt Tamborine Village [-27 53, 153 8], 15/i/57, BBL. NEW SOUTH WALES:
<emphasis box="[1001,1177,694,717]" italics="true" pageId="21" pageNumber="514">
<taxonomicName box="[1001,1177,694,717]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">M. impaternata</taxonomicName>
</emphasis>
was found common at localities near Armidale [-30 30, 151 40], NSW, in Dec/Jan. 199596 and Nov. 1999 by JACP collectors. It is sympatric there with
<emphasis box="[415,574,765,789]" italics="true" pageId="21" pageNumber="514">
<taxonomicName box="[415,570,765,789]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="croslandi">M. croslandi</taxonomicName>
.
</emphasis>
Localities elsewhere include: Corang River Bridge [-35 12, 150 03]; Charleyong [-35 15, 149 55], HI87156; Yowrie [-36 19, 149 44], HI87161, HI87162. AUSTRALIAN CAPITAL TERRITORY: Canberra [-35 18, 149 8], HI85372, 373, HI87237. AAVAS collections are from the Australian National Botanic Gardens site [-35 17, 149 7], and the suburbs of Cook [-35 16, 149 4] and Hawker [-35 15, 149 2], VICTORIA: Mayfield [-35 12, 149.48], HI87155.
</paragraph>
<paragraph pageId="21" pageNumber="514">
<emphasis box="[199,457,946,970]" italics="true" pageId="21" pageNumber="514">
<taxonomicName box="[199,457,946,970]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">Myrmecia impaternata</taxonomicName>
</emphasis>
(along with the locally more frequent
<emphasis box="[893,1036,945,970]" italics="true" pageId="21" pageNumber="514">
<taxonomicName box="[893,1036,945,970]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="croslandi">M. croslandi</taxonomicName>
</emphasis>
) is common in urban and suburban Canberra (see below).
</paragraph>
</subSubSection>
<subSubSection pageId="21" pageNumber="514" type="diagnosis">
<paragraph pageId="21" pageNumber="514">
<emphasis bold="true" box="[199,416,1017,1042]" pageId="21" pageNumber="514">Worker diagnosis.</emphasis>
General features as illustrated and in key couplets 1 4 above, which cover leg-coloration. The brassy color of the cephalic pubescence can be hard to discern stereomicroscopically using some types of illumination lamp, and color temperatures of the light provided. It is best observed beyond the near-side eye in acute diagonal lateral view of the head. This pubescence is arguably a legacy of the
<taxonomicName box="[1117,1229,1125,1149]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="banksi">
<emphasis box="[1117,1229,1125,1149]" italics="true" pageId="21" pageNumber="514">M. banksi</emphasis>
</taxonomicName>
hybrid parentage. Otherwise
<taxonomicName box="[277,455,1162,1185]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[277,455,1162,1185]" italics="true" pageId="21" pageNumber="514">M. impaternata</emphasis>
</taxonomicName>
is similar in physiognamy and sculpturation to small/mediumsized workers of the second parental species,
<taxonomicName box="[435,564,1197,1221]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="pilosula">
<emphasis box="[435,564,1197,1221]" italics="true" pageId="21" pageNumber="514">M. pilosula</emphasis>
</taxonomicName>
(Eastern Race). There is some size-related graded variation in cephalic and dorsal mesosomal sculpturation between small and large specimens, somewhat as in both races of
<taxonomicName box="[1264,1392,1233,1258]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="pilosula">
<emphasis box="[1264,1392,1233,1258]" italics="true" pageId="21" pageNumber="514">M. pilosula</emphasis>
</taxonomicName>
but less extreme, especially at the high, more intensively-sculptured end of the range.
</paragraph>
<paragraph pageId="21" pageNumber="514">
For significant scientific reasons discussed below investigation of the comparative biology of, and possible reproductive relationships between,
<taxonomicName box="[557,731,1342,1366]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[557,731,1342,1366]" italics="true" pageId="21" pageNumber="514">M. impaternata</emphasis>
</taxonomicName>
and
<taxonomicName box="[786,928,1341,1366]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="croslandi">
<emphasis box="[786,928,1341,1366]" italics="true" pageId="21" pageNumber="514">M. croslandi</emphasis>
</taxonomicName>
is a prime scientific subject. The two species may be readily differentiated using the characters of couplet 2 of the key to species above.
</paragraph>
</subSubSection>
<subSubSection pageId="21" pageNumber="514" type="description">
<paragraph pageId="21" pageNumber="514">
<emphasis bold="true" box="[199,345,1413,1438]" pageId="21" pageNumber="514">Dimensions.</emphasis>
(Holotype, smallest paratype, largest paratype (mm): TL = 13.01, 10.87, 13.04; HW = 2.47, 2.22, 2.49; HL = 2.27, 206, 2.27; CI = 108, 107, 109; EL = 0.96, 0.88, 0.97; OI = 39, 39, 39; SL = 1.87, 1.84, 1.94; SI = 76, 82, 78; PW = 1.58, 1.42, 1.59; WL = 3.52, 3.34, 3.75; PetW = 0.97, 0.83, 0.96; PpetW = 1.39, 1.22, 1.33.
</paragraph>
</subSubSection>
<subSubSection pageId="21" pageNumber="514" type="etymology">
<paragraph pageId="21" pageNumber="514">
<emphasis bold="true" box="[199,333,1522,1545]" pageId="21" pageNumber="514">Etymology.</emphasis>
The name
<taxonomicName box="[460,597,1522,1545]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[460,597,1522,1545]" italics="true" pageId="21" pageNumber="514">impaternata</emphasis>
</taxonomicName>
is based on the Latinate (not truly Latin) biological term “impaternate” (= fatherless as a result of parthenogenesis). For explanation see the section on reproductive biology below.
</paragraph>
</subSubSection>
<subSubSection pageId="21" pageNumber="514" type="description">
<paragraph pageId="21" pageNumber="514">
<emphasis bold="true" box="[199,330,1594,1617]" pageId="21" pageNumber="514">Karyology.</emphasis>
Details are provided by Imai, Taylor et. al. (1994), and by Taylor, Imai and Hasegawa (in preparation).
<taxonomicName box="[307,570,1630,1654]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[307,570,1630,1654]" italics="true" pageId="21" pageNumber="514">Myrmecia impaternata</emphasis>
</taxonomicName>
has an allodiploid karyotype: n=5 or 14, 2n=19. The 5-chromosome set closely matches one of the haploid sets of
<taxonomicName box="[624,734,1665,1690]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="banksi">
<emphasis box="[624,734,1665,1690]" italics="true" pageId="21" pageNumber="514">M. banksi</emphasis>
</taxonomicName>
, while the 14-chromosome haploid set is generally matched in the Eastern Race of
<taxonomicName box="[384,514,1701,1726]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="pilosula">
<emphasis box="[384,514,1701,1726]" italics="true" pageId="21" pageNumber="514">M. pilosula</emphasis>
</taxonomicName>
most closely resembling chromosomes from a colony (HI87130) collected at Wambrook Creek (36º11'S, 148º56), near Cooma, NSW. On these grounds
<taxonomicName box="[1026,1140,1737,1762]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="banksi">
<emphasis box="[1026,1140,1737,1762]" italics="true" pageId="21" pageNumber="514">M. banksi</emphasis>
</taxonomicName>
and
<taxonomicName box="[1201,1332,1737,1762]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="pilosula">
<emphasis box="[1201,1332,1737,1762]" italics="true" pageId="21" pageNumber="514">M. pilosula</emphasis>
</taxonomicName>
(Eastern Race) (or close ancestral stocks) are identified here as the parental species which hybridized to originate
<taxonomicName class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis italics="true" pageId="21" pageNumber="514">M. impaternata</emphasis>
</taxonomicName>
. See also Imai, Taylor
<emphasis box="[546,603,1809,1834]" italics="true" pageId="21" pageNumber="514">et al.</emphasis>
(1994) appendix and fig. 8 (p. 150).
</paragraph>
</subSubSection>
<subSubSection lastPageId="23" lastPageNumber="516" pageId="21" pageNumber="514" type="biology_ecology">
<paragraph pageId="21" pageNumber="514">
<emphasis bold="true" box="[199,417,1845,1870]" pageId="21" pageNumber="514">Field associations.</emphasis>
All
<taxonomicName box="[469,645,1846,1870]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[469,645,1846,1870]" italics="true" pageId="21" pageNumber="514">M. impaternata</emphasis>
</taxonomicName>
sympatric associations throughout its known distribution are with
<taxonomicName class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="21" pageNumber="514" phylum="Arthropoda" rank="species" species="croslandi">
<emphasis italics="true" pageId="21" pageNumber="514">M. croslandi</emphasis>
</taxonomicName>
. Nests of both species are frequently encountered interspersed in Canberra parks and gardens, suburban roadside grass lawn “nature strips” and in local grassy bushland. Colonies of the two have often been encountered by the author and other researchers only a few meters apart.
</paragraph>
<caption pageId="22" pageNumber="515" targetBox="[282,1304,202,1933]" targetPageId="22">
<paragraph pageId="22" pageNumber="515">
<emphasis bold="true" box="[151,339,1956,1977]" pageId="22" pageNumber="515">FIGURES 1618.</emphasis>
<taxonomicName box="[345,577,1956,1977]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="22" pageNumber="515" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[345,577,1956,1977]" italics="true" pageId="22" pageNumber="515">Myrmecia impaternata</emphasis>
</taxonomicName>
, National Botanic Gardens, Canberra, ACT. Standard views. HW 2.29 mm, WL 3.35 mm, PW 1.42 mm.
</paragraph>
</caption>
<paragraph pageId="23" pageNumber="516">
<emphasis bold="true" box="[199,466,151,176]" pageId="23" pageNumber="516">Reproductive biology.</emphasis>
There has been persistent historical failure by JACP researchers, and the author subsequently, to discover males in
<emphasis box="[585,768,189,212]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[585,768,189,212]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">M. impaternata</taxonomicName>
</emphasis>
nests, despite targeted excavation of field colonies at appropriate seasons over many years. Only two male-right colonies have ever been located and collected. Both were closely adjacent at the Canberra Botanic Gardens
</paragraph>
</subSubSection>
<subSubSection pageId="23" pageNumber="516" type="discussion">
<paragraph pageId="23" pageNumber="516">
Research on males and queens from these nests by Taylor, Imai and Hasegawa (to be published elsewhere) has investigated the reproductive biology of
<emphasis box="[620,797,333,356]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[620,797,333,356]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">M. impaternata</taxonomicName>
</emphasis>
. It convincingly demonstrates that
<taxonomicName box="[1206,1385,333,356]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">
M.
<emphasis box="[1248,1385,333,356]" italics="true" pageId="23" pageNumber="516">impaternata</emphasis>
</taxonomicName>
is a sperm-dependent gynogenetic taxon in which unreduced eggs require contact with sperm or spermatic fluid (specifically without the occurrence of fertilization) in order to develop parthenogenetically, and thus to produce diploid workers and gynes (see Kokko
<emphasis box="[592,642,439,464]" italics="true" pageId="23" pageNumber="516">et al</emphasis>
. (2008) for theoretical background). The necessary spermatic material is evidentially obtained by gynes through copulation with nominally conspecific donor males bred in
<emphasis box="[1299,1436,477,500]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[1299,1436,477,500]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">impaternata</taxonomicName>
</emphasis>
nests. In this example the sperm cells dissected from male testes and gyne spermathecae were identically and characteristically structurally degenerate and putatively incapable of actually effecting fertilization. Their presence in both sexes importantly attests previous mating between relevant males and gynes. The wheres and whens of copulation are not known.
</paragraph>
<paragraph pageId="23" pageNumber="516">In other known sperm-dependent gynogenetic animals (all of which are hybrid-originated allodiploid entities, including various fish and amphibian species, none of which are known to possess a male sex) sperm is obtained for this unusual purpose by the females through parasitic copulation with males of other separate, sympatric, congeneric and usually closely related donor species. The distribution, stable presence and long-term population survival of such gynogenetic taxa is totally dependent on sympatric associations with sperm-donor host species, a factor which critically restrains their distributional ranges and dispersability, but ultimately ensures their survival as species in nature.</paragraph>
<paragraph pageId="23" pageNumber="516">
Because males in ants are genetically haploid, it is suggested that those produced by
<emphasis box="[1154,1291,909,932]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[1154,1291,909,932]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">impaternata</taxonomicName>
</emphasis>
females will likely be of two types, genetically, karyologically and perhaps morphologically equivalent to males of the putative parental species
<emphasis box="[339,451,980,1004]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[339,451,980,1004]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="banksi">M. banksi</taxonomicName>
</emphasis>
and
<emphasis box="[509,638,980,1004]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[509,638,980,1004]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="pilosula">M. pilosula</taxonomicName>
</emphasis>
(Eastern Race). Remarkably, males produced in
<emphasis box="[1197,1334,981,1004]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[1197,1334,981,1004]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">impaternata</taxonomicName>
</emphasis>
colonies would technically therefore not be conspecific with their
<taxonomicName box="[847,984,1016,1040]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[847,984,1016,1040]" italics="true" pageId="23" pageNumber="516">impaternata</emphasis>
</taxonomicName>
mothers. Taylor, Imai and Hasegawa conclude that: “
<taxonomicName box="[328,503,1053,1076]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[328,503,1053,1076]" italics="true" pageId="23" pageNumber="516">M. impaternata</emphasis>
</taxonomicName>
thus has no need to maintain risky, restrictive parasitic affiliation or sympatry with other free-living, closely-related sperm-donor host species. It is apparently able to produce the necessary allospecific males by accessing its own genome!” The authors also suggest that
<taxonomicName box="[1070,1246,1125,1148]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[1070,1246,1125,1148]" italics="true" pageId="23" pageNumber="516">M. impaternata</emphasis>
</taxonomicName>
queens might at times more usually operate as sperm parasites of
<taxonomicName box="[711,854,1159,1184]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="croslandi">
<emphasis box="[711,854,1159,1184]" italics="true" pageId="23" pageNumber="516">M. croslandi</emphasis>
</taxonomicName>
by obtaining sperm allospecifically from
<taxonomicName box="[1331,1436,1159,1184]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="croslandi">
<emphasis box="[1331,1436,1159,1184]" italics="true" pageId="23" pageNumber="516">croslandi</emphasis>
</taxonomicName>
males. This hypothesis is encouraged by the persistent historical failure to discover males in
<taxonomicName box="[1189,1363,1197,1220]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[1189,1363,1197,1220]" italics="true" pageId="23" pageNumber="516">M. impaternata</emphasis>
</taxonomicName>
nests.
</paragraph>
<paragraph pageId="23" pageNumber="516">
<emphasis bold="true" box="[199,593,1231,1256]" pageId="23" pageNumber="516">Status as a taxonomic species.</emphasis>
Despite its apparent hybrid origin and reproduction by theletokous parthenogenesis,
<taxonomicName box="[349,528,1269,1292]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[349,528,1269,1292]" italics="true" pageId="23" pageNumber="516">M. impaternata</emphasis>
</taxonomicName>
functions in nature as a biological species. It is accorded specific taxonomic status here following the precepts of Maslin (1968) and Cole (1985), on the grounds that it is a genetically and historically unique, self perpetuating, separately evolving entity, reproductively isolated from its ancestors and sympatrically-associated related species.
</paragraph>
<paragraph pageId="23" pageNumber="516">
Taylor, Imai &amp; Hasegawa raise the possibility that several
<emphasis box="[928,1119,1413,1436]" italics="true" pageId="23" pageNumber="516">M. impaternata-</emphasis>
related hybrid clones with separately evolved alternative reproductive arrangements could be present in nature, each derived from a different foundation hybridization event between
<taxonomicName box="[614,727,1484,1508]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="banksi">
<emphasis box="[614,727,1484,1508]" italics="true" pageId="23" pageNumber="516">M. banksi</emphasis>
</taxonomicName>
and
<emphasis box="[787,917,1484,1508]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[787,917,1484,1508]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="pilosula">M. pilosula</taxonomicName>
</emphasis>
(Eastern Race). Note that Imai, Taylor
<emphasis box="[1377,1437,1484,1508]" italics="true" pageId="23" pageNumber="516">et al.</emphasis>
(1994) recognized two forms of
<emphasis box="[525,702,1521,1544]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[525,702,1521,1544]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">M. impaternata</taxonomicName>
</emphasis>
(PBF1-1 and PBF1-2) differing in details of leg coloration (see their fig 10). These could arguably represent separately originated
<taxonomicName box="[922,995,1556,1580]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="banksi">
<emphasis box="[922,995,1556,1580]" italics="true" pageId="23" pageNumber="516">banksi</emphasis>
</taxonomicName>
x
<emphasis box="[1029,1119,1556,1580]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[1029,1119,1556,1580]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="pilosula">pilosula</taxonomicName>
</emphasis>
hybrid entities. Alternative hybrid lineages, if demonstrated, could not be considered formally conspecific with one another following the precepts of Maslin (1968) and Cole (1985) discussed above, nor would they be.
<emphasis box="[1043,1300,1629,1652]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[1043,1300,1629,1652]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">Myrmecia impaternata</taxonomicName>
</emphasis>
and its kind could well have an interesting scientific future!
</paragraph>
<paragraph pageId="23" pageNumber="516">
<emphasis bold="true" box="[199,436,1701,1724]" pageId="23" pageNumber="516">Research prospects.</emphasis>
Further understanding of the reproductive biology of
<emphasis box="[1037,1212,1701,1724]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[1037,1212,1701,1724]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">M. impaternata</taxonomicName>
</emphasis>
is greatly desirable. Of particular interest are: (1) determining whether sperm or spermatic materials actually enter the egg cytoplasm or not; (2) investigation of the possibility that two classes of males (comparable respectively to those found in colonies of
<emphasis box="[280,390,1807,1832]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[280,390,1807,1832]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="banksi">M. banksi</taxonomicName>
</emphasis>
and
<emphasis box="[443,570,1807,1832]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[443,570,1807,1832]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="pilosula">M. pilosula</taxonomicName>
</emphasis>
[Eastern Race]) are developed from haploid
<emphasis box="[1070,1207,1808,1832]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[1070,1207,1808,1832]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">impaternata</taxonomicName>
</emphasis>
queen-laid eggs; (3) testing the possibility that production of “
<emphasis box="[637,774,1845,1868]" italics="true" pageId="23" pageNumber="516">
<taxonomicName box="[637,774,1845,1868]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">impaternata</taxonomicName>
</emphasis>
” males and their presence in nests might in this case be unusual; (4) finding whether
<taxonomicName box="[484,621,1880,1904]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="impaternata">
<emphasis box="[484,621,1880,1904]" italics="true" pageId="23" pageNumber="516">impaternata</emphasis>
</taxonomicName>
might also (more usually?) maintain a parasitic copulatory relationship with
<taxonomicName box="[210,355,1916,1940]" class="Insecta" family="Formicidae" genus="Myrmecia" higherTaxonomySource="CoL" kingdom="Animalia" order="Hymenoptera" pageId="23" pageNumber="516" phylum="Arthropoda" rank="species" species="croslandi">
<emphasis box="[210,355,1916,1940]" italics="true" pageId="23" pageNumber="516">M. croslandi</emphasis>
</taxonomicName>
, considering their frequent, very proximate, and wide-ranging sympatric co-presence; and (5) determining when and where copulation occurs in either of these scenarios.
</paragraph>
</subSubSection>
</treatment>
</document>